The human brain displays rich communication dynamics that are thought to be particularly well-reflected in its marked community structure. Yet, the precise relationship between community structure in structural brain networks and the communication dynamics that can emerge therefrom is not well-understood. In addition to offering insight into the structure-function relationship of networked systems, such an understanding is a critical step towards the ability to manipulate the brain's large-scale dynamical activity in a targeted manner. We investigate the role of community structure in the controllability of structural brain networks. At the region level, we find that certain network measures of community structure are sometimes statistically correlated with measures of linear controllability. However, we then demonstrate that this relationship depends on the distribution of network edge weights. We highlight the complexity of the relationship between community structure and controllability by performing numerical simulations using canonical graph models with varying mesoscale architectures and edge weight distributions. Finally, we demonstrate that weighted subgraph centrality, a measure rooted in the graph spectrum, and which captures higher-order graph architecture, is a stronger and more consistent predictor of controllability. Our study contributes to an understanding of how the brain's diverse mesoscale structure supports transient communication dynamics.
Modularity is an important topological attribute for functional brain networks. Recent studies have reported that modularity of functional networks varies not only across individuals being related to demographics and cognitive performance, but also within individuals co-occurring with fluctuations in network properties of functional connectivity, estimated over short time intervals. However, characteristics of these time-resolved functional networks during periods of high and low modularity have remained largely unexplored. In this study we investigate spatiotemporal properties of time-resolve…
In cognitive network neuroscience, the connectivity and community structure of the brain network is related to cognition. Much of this research has focused on two measures of connectivity - modularity and flexibility - which frequently have been examined in isolation. By using resting state fMRI data from 52 young adults, we investigate the relationship between modularity, flexibility and performance on cognitive tasks. We show that flexibility and modularity are highly negatively correlated. However, we also demonstrate that flexibility and modularity make unique contributions to explain task…
Over one third of the estimated 3 million people with epilepsy in the US are medication resistant. Responsive neurostimulation from chronically implanted electrodes provides a promising treatment option and alternative to resective surgery. However, determining personalized optimal stimulation parameters, including when and where to intervene to guarantee a positive patient outcome, is a major open challenge. Network neuroscience and control theory offer useful tools that may guide improvements in parameter selection for control of anomalous neural activity. Here we use a novel method to chara…
We investigate the relationship of resting-state fMRI functional connectivity estimated over long periods of time with time-varying functional connectivity estimated over shorter time intervals. We show that using Pearson's correlation to estimate functional connectivity implies that the range of fluctuations of functional connections over short time scales is subject to statistical constraints imposed by their connectivity strength over longer scales. We present a method for estimating time-varying functional connectivity that is designed to mitigate this issue and allows us to identify episo…
Neuroimaging data can be represented as networks of nodes and edges that capture the topological organization of the brain connectivity. Graph theory provides a general and powerful framework to study these networks and their structure at various scales. By way of example, community detection methods have been widely applied to investigate the modular structure of many natural networks, including brain functional connectivity networks. Sparsification procedures are often applied to remove the weakest edges, which are the most affected by experimental noise, and to reduce the density of the gra…
The human brain network is modular--comprised of communities of tightly interconnected nodes. This network contains local hubs, which have many connections within their own communities, and connector hubs, which have connections diversely distributed across communities. A mechanistic understanding of these hubs and how they support cognition has not been demonstrated. Here, we leveraged individual differences in hub connectivity and cognition. We show that a model of hub connectivity accurately predicts the cognitive performance of 476 individuals in four distinct tasks. Moreover, there is a g…
This paper studies the link between resting-state functional connectivity (FC), measured by the correlations of the fMRI BOLD time courses, and structural connectivity (SC), estimated through fiber tractography. Instead of a static analysis based on the correlation between SC and the FC averaged over the entire fMRI time series, we propose a dynamic analysis, based on the time evolution of the correlation between SC and a suitably windowed FC. Assessing the statistical significance of the time series against random phase permutations, our data show a pronounced peak of significance for time wi…
Network modularity is a key feature for efficient information processing in the human brain. This information processing is however dynamic and networks can reconfigure at very short time period, few hundreds of millisecond. This requires neuroimaging techniques with sufficient time resolution. Here we use the dense electroencephalography, EEG, source connectivity methods to identify cortical networks with excellent time resolution, in the order of millisecond. We identify functional networks during picture naming task. Two categories of visual stimuli were presented, meaningful (tools, animal…
Music listening involves many simultaneous neural operations, including auditory processing, working memory, temporal sequencing, pitch tracking, anticipation, reward, and emotion, and thus, a full investigation of music cognition would benefit from whole-brain analyses. Here, we quantify whole-brain activity while participants listen to a variety of music and speech auditory pieces using two network measures that are grounded in complex systems theory: modularity, which measures the degree to which brain regions are interacting in communities, and flexibility, which measures the rate that bra…
The psychosis spectrum is associated with structural dysconnectivity concentrated in transmodal association cortex. However, understanding of this pathophysiology has been limited by an exclusive focus on the direct connections to a region. Using Network Control Theory, we measured variation in both direct and indirect structural connections to a region to gain new insights into the pathophysiology of the psychosis spectrum. We used psychosis symptom data and structural connectivity in 1,068 youths aged 8 to 22 years from the Philadelphia Neurodevelopmental Cohort. Applying a Network Control T…
The large-scale structural ingredients of the brain and neural connectomes have been identified in recent years. These are, similar to the features found in many other real networks: the arrangement of brain regions into modules and the presence of highly connected regions (hubs) forming rich-clubs. Here, we examine how modules and hubs shape the collective dynamics on networks and we find that both ingredients lead to the emergence of complex dynamics. Comparing the connectomes of C. elegans, cats, macaques and humans to surrogate networks in which either modules or hubs are destroyed, we fin…
Predicting how the brain can be driven to specific states by means of internal or external control requires a fundamental understanding of the relationship between neural connectivity and activity. Network control theory is a powerful tool from the physical and engineering sciences that can provide insights regarding that relationship; it formalizes the study of how the dynamics of a complex system can arise from its underlying structure of interconnected units. Given the recent use of network control theory in neuroscience, it is now timely to offer a practical guide to methodological conside…
Network control theory has recently emerged as a promising approach for understanding brain function and dynamics. By operationalizing notions of control theory for brain networks, it offers a fundamental explanation for how brain dynamics may be regulated by structural connectivity. While powerful, the approach does not currently consider other non-structural explanations of brain dynamics. Here we extend the analysis of network controllability by formalizing the evolution of neural signals as a function of effective inter-regional coupling and pairwise signal covariance. We find that functio…
Understanding the complex hierarchical topology of functional brain networks is a key aspect of functional connectivity research. Such topics are obscured by the widespread use of sparse binary network models which are fundamentally different to the complete weighted networks derived from functional connectivity. We introduce two techniques to probe the hierarchical complexity of topologies. Firstly, a new metric to measure hierarchical complexity; secondly, a Weighted Complex Hierarchy (WCH) model. To thoroughly evaluate our techniques, we generalise sparse binary network archetypes to weight…
As the human brain develops, it increasingly supports coordinated control of neural activity. The mechanism by which white matter evolves to support this coordination is not well understood. We use a network representation of diffusion imaging data from 882 youth ages 8 to 22 to show that white matter connectivity becomes increasingly optimized for a diverse range of predicted dynamics in development. Notably, stable controllers in subcortical areas are negatively related to cognitive performance. Investigating structural mechanisms supporting these changes, we simulate network evolution with …
Emerging evidence shows that the modular organization of the human brain allows for better and efficient cognitive performance. Many of these cognitive functions are very fast and occur in subsecond time scale such as the visual object recognition. Here, we investigate brain network modularity while controlling stimuli meaningfulness and measuring participant reaction time. We particularly raised two questions: i) does the dynamic brain network modularity change during the recognition of meaningful and meaningless visual images? And ii) is there a correlation between network modularity and the…
Modularity plays an important role in brain networks' architecture and influences its dynamics and the ability to integrate and segregate different modules of cerebral regions. Alterations in community structure are associated with several clinical disorders, specially schizophrenia, although its time evolution is not clear yet. In the present work, we analyze fMRI functional networks of $65$ healthy subjects (HC) and $44$ patients of schizophrenia (SZ), $28$ of them in a chronic state (CR) of illness, and $16$ at early stage (ES). We find clear differences in edges' weights distribution, netw…
In this paper we utilized the concept of stable phase synchronization topography - synchrostates - over the scalp derived from EEG recording for formulating brain connectivity network in Autism Spectrum Disorder (ASD) and typically-growing children. A synchronization index is adapted for forming the edges of the connectivity graph capturing the stability of each of the synchrostates. Such network is formed for 11 ASD and 12 control group children. Comparative analyses of these networks using graph theoretic measures show that children with autism have a different modularity of such networks fr…
The connectome, or the entire connectivity of a neural system represented by network, ranges various scales from synaptic connections between individual neurons to fibre tract connections between brain regions. Although the modularity they commonly show has been extensively studied, it is unclear whether connection specificity of such networks can already be fully explained by the modularity alone. To answer this question, we study two networks, the neuronal network of C. elegans and the fibre tract network of human brains yielded through diffusion spectrum imaging (DSI). We compare them to th…
Recently we developed a novel method for assessing the hierarchical modularity of functional brain networks - the probability associated community estimation(PACE). The PACE algorithm is unique in that it permits a dual formulation, thus yielding equivalent connectome modular structure regardless of whether considering positive or negative edges. This method was rigorously validated using F1000 and HCP data. We detected novel sex differences in resting-state connectivity that were not previously reported. This current study more thoroughly examined sex differences as a function of age and thei…
Background: A therapeutic intervention in psychiatry can be viewed as an attempt to influence the brain's large-scale, dynamic network state transitions underlying cognition and behavior. Building on connectome-based graph analysis and control theory, Network Control Theory is emerging as a powerful tool to quantify network controllability - i.e., the influence of one brain region over others regarding dynamic network state transitions. If and how network controllability is related to mental health remains elusive. Methods: From Diffusion Tensor Imaging data, we inferred structural connectivit…
Community detection algorithms have been widely used to study the organization of complex systems like the brain. A principal appeal of these techniques is their ability to identify a partition of brain regions (or nodes) into communities, where nodes within a community are densely interconnected. In their simplest application, community detection algorithms are agnostic to the presence of community hierarchies, but a common characteristic of many neural systems is a nested hierarchy. To address this limitation, we exercise a multi-scale extension of a community detection technique known as mo…
Cognition is supported by neurophysiological processes that occur both in local anatomical neighborhoods and in distributed large-scale circuits. Recent evidence from network control theory suggests that white matter pathways linking large-scale brain regions provide a critical substrate constraining the ability of single areas to affect control on those processes. Yet, no direct evidence exists for a relationship between brain network controllability and cognitive control performance. Here, we address this gap by constructing structural brain networks from diffusion tensor imaging data acquir…
Circadian rhythms synchronize a variety of physiological processes ranging from neural activity and hormone secretion to sleep cycles and feeding habits. Despite significant diurnal variation, time-of-day (TOD) is rarely recorded or analyzed in human brain research. Moreover, sleep-wake patterns, diurnal preferences, and daytime alertness vary across individuals, known as sleep chronotypes. Here, we performed graph-theory network analysis on resting-state functional MRI (rs-fMRI) data to explore topological differences in whole-brain functional networks between morning and evening sessions (TO…
The relationship of network structure and dynamics is one of most extensively investigated problems in the theory of complex systems of the last years. Understanding this relationship is of relevance to a range of disciplines -- from Neuroscience to Geomorphology. A major strategy of investigating this relationship is the quantitative comparison of a representation of network architecture (structural connectivity) with a (network) representation of the dynamics (functional connectivity). Analysing such SC/FC relationships has over the past years contributed substantially to our understanding o…
Hierarchical Bayesian inference for community detection and connectivity of functional brain networks
ShanghaiTech University（上海科技大学）；Monash University（蒙纳士大学）；The Hong Kong Polytechnic University（香港理工大学）；University College London（伦敦大学学院）；CIFAR（加拿大高级研究所）
Functional connectivity refers to the temporal statistical relationship between spatially distinct brain regions and is usually inferred from the time series coherence/correlation in brain activity between regions of interest. In human functional brain networks, the network structure is often inferred from functional magnetic resonance imaging (fMRI) blood oxygen level dependent (BOLD) signal. Since the BOLD signal is a proxy for neuronal activity, it is of interest to learn the latent functional network structure. Additionally, despite a core set of observations about functional networks such…
Artificial and natural neural network models are a new toolkit which could be potentially have been used for clarifying of complex brain functions. To attend this goal, such models need to be neurobiologically realistic. However, although neural networks have advanced keenly in recent decades their strict similarity in aspects of brain anatomy and physiology is imperfect. In this work we discuss different types of neural models, including localist, attractor and deep network models, and also identify aspects under which their biological credibility can be improved. These conditions range from …
Monash University Malaysia（马来西亚蒙纳士大学）；King Abdullah University of Science and Technology（阿卜杜拉国王科技大学）；Xiamen University Malaysia（厦门大学马来西亚分校）
We present a unified statistical framework for characterizing community structure of brain functional networks that captures variation across individuals and evolution over time. Existing methods for community detection focus only on single-subject analysis of dynamic networks; while recent extensions to multiple-subjects analysis are limited to static networks. To overcome these limitations, we propose a multi-subject, Markov-switching stochastic block model (MSS-SBM) to identify state-related changes in brain community organization over a group of individuals. We first formulate a multilayer…
A major goal shared by neuroscience and collective behavior is to understand how dynamic interactions between individual elements give rise to behaviors in populations of neurons and animals, respectively. This goal has recently become within reach thanks to techniques providing access to the connectivity and activity of neuronal ensembles as well as to behaviors among animal collectives. The next challenge using these datasets is to unravel network mechanisms generating population behaviors. This is aided by network theory, a field that studies structure-function relationships in interconnect…
The understanding of neural activity patterns is fundamentally linked to an understanding of how the brain's network architecture shapes dynamical processes. Established approaches rely mostly on deviations of a given network from certain classes of random graphs. Hypotheses about the supposed role of prominent topological features (for instance, the roles of modularity, network motifs, or hierarchical network organization) are derived from these deviations. An alternative strategy could be to study deviations of network architectures from regular graphs (rings, lattices) and consider the impl…
Hierarchical Multiscale Structure-Function Coupling for Brain Connectome Integration
School of Medicine, University of Dundee, UK（邓迪大学医学院）；School of Science and Engineering, University of Dundee, UK（邓迪大学科学与工程学院）；College of Medicine and Biological Information Engineering, Northeastern University, China（东北大学医学院与生物信息工程学院）；College of Medical Informatics, Chongqing Medical University, China（重庆医科大学医学信息学院）；School of Airspace Science and Engineering, Shandong University, China（山东大学航空航天科学与工程学院）；Chinese Institute for Brain Research, China（中国脑科学研究院）；Department of Applied Mathematics and Theoretical Physics, University of Cambridge, UK（剑桥大学应用数学与理论物理系）；Department of Clinical Neurosciences, University of Cambridge, UK（剑桥大学临床神经科学系）
The primate heteromodal cortex presents an evident functional modularity at a mesoscopic level, with physiological and anatomical evidence pointing to it as likely substrate of long-term memory. In order to investigate some of its properties, a model of multimodular autoassociator is studied. Each of the many modules represents a neocortical functional ensemble of recurrently connected neurons and operates as a Hebbian autoassociator, storing a number of local features which it can recall upon cue. The global memory patterns are made of combinations of features sparsely distributed across the …
Stimulation-based control of dynamic brain networks
University of Pennsylvania（宾夕法尼亚大学）；US Army Research Laboratory（美国陆军研究实验室）；University at Buffalo, SUNY（纽约州立大学布法罗分校）；University of California, Riverside（加州大学河滨分校）；University of California, Santa Barbara（加州大学圣塔芭芭拉分校）
